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​The combtooth blennies (f. Blenniidae) is a diverse family of primarily
marine fishes with approximately 387 species that inhabit subtidal,
intertidal, supralittoral habitats in tropical and warm temperate
regions throughout the world. The Blenniidae has typically been divided
into six groups based on morphological characters: Blenniini, Nemophini,
Omobranchini, Phenablenniini, Parablenniini, and Salariini. There is,
however, considerable debate over the validity of these groups and their
relationships. Since little is known about the relationships in this
group, other aspects of their evolutionary history, such as habitat
evolution and remain unexplored. Herein, we use Bayesian and maximum
likelihood analyses of four nuclear loci (ENC1, myh6, ptr, and tbr1)
from 102 species, representing 41 genera, to resolve the phylogeny of
the Blenniidae, determine the validity of the previously recognized
groupings, and explore the evolution of habitat association using
ancestral state reconstruction. Bayesian and maximum likelihood analyses
of the resulting 3100 bp of DNA sequence produced nearly identical
topologies, and identified many well-supported clades. Of these clades,
Nemophini was the only traditionally recognized group strongly supported
as monophyletic. This highly resolved and thoroughly sampled blenniid
phylogeny provides strong evidence that the traditional rank-based
classification does not adequately delimit monophyletic groups with the
Blenniidae. This phylogeny redefines the taxonomy of the group and
supports the use of 13 unranked clades for the classification of
blenniids. Ancestral state reconstructions identified four independent
invasions of intertidal habitats within the Blenniidae, and subsequent
invasions into supralittoral and freshwater habitats from these groups.
The independent invasions of intertidal habitats are likely to have
played an important role in the evolutionary history of blennies.